Records of the Western Australian Museum 17: 213-220 (1995).

Morphological variation in Bearded Tomb Bats (Taphozous) in Tenggara and Nusa Tenggara Timur,

D.J. Kitchenerl and A. Suyanto2

1 Western Australian Museum, Francis Street, Perth, Western 6000 2 Balitbang Zoologi, LlPI, Jalan Ir. H. Juanda 9, Bogor, Indonesia 16122

Abstract - Bearded Tomb Bats, TapholollS melanopogoll and T acllates. were recently collected on , Tanimbar and Kai , eastern Indonesia. Morphological comparisons of skull, dentary, dental and external characters and a univariate and multivariate statistical analyses of 138 specimens, indicated that the animals from Wetar were T a. achates. Those from the Tanimbar and were morphologically distinct from both T. melanopogon and T. a. achates, but were closer to the latter taxon. They are herein described as a new subspecies of T. achates, T. achates minor

INTRODUCTION more isolated from the other islands in these areas An earlier study of the morphological variation (Kitchener and Maryanto 1993; Kitchener et al. of Bearded Tomb Bats in Nusa Tenggara, 1993b; 1995a,b). Indonesia, by Kitchener et al. (1993a) recognised This paper will describe the morphological the occurrence in that region of two species, variation of the Bearded Tomb Bats from the Taphozous m. meIanopogon Temminck, 1841 and T. southern region of Maluku Tenggara and report on achates Thomas, 1915. The former was found on the taxonomic status of these bats. islands in the Inner (Lombok, , Moyo and Alor), as well as in the Outer Banda Arc. Koopman and Cordon (1992) MATERIAL AND METHODS also report that a specimen of T. meIanopogon exists Measurements from 111 adult specimens used in in the American Museum of Natural History from the previous analysis of morphological variation in Kei (= Kai) Island. Taphozous achates occurred only Taphozous in this region presented in Kitchener et on islands in the Outer Banda Arc (Savu, Roti and al. (1993a) were combined with measurements Semau and possibly also on Timor). from an additional 33 adult specimens from Wetar Kitchener et al. (1993a) noted that while there was Island (9), Kai Besar Island (12) and Selaru Island, little morphological differentiation in T. m. Tanimbar Croup (12) for statistical analysis. The melanopogon among islands in Nusa Tenggara, measurements recorded were the same as those there appeared to be some differentiation among documented in that previous study. However, populations of T. achates. These authors suggest palate length in both this and the previous study that T. m. meIanopogon is Laurasian in origin and was measured from the posterior margin of the may be a relatively recent inhabitant of Nusa palate to the posterior margin of the anterior Tenggara and speculate that '7. achates alone will palatal incisor, and not to the anterior edge of the be found to occur on the smaller Condwanic Cl alveoli as figured in Kitchener et al. (1993a: islands such as , Babar, Tanimbar, Kei (= Kai) figure 1). and Aru". The recent collection of Bearded Tomb The measurements were as follows: CSL, greatest Bats from Wetar, Tanimbar and Kai Islands by staff skull length; IOW, least interorbital breadth; POW, from the Western Australian Museum and postorbital breadth; CW, cranial breadth; MW, Museum Zoologicurn Bogoriense allows for this mastoid width; ZW, zygomatic width; PPL, post speculation to be examined. palatal length; PL, palatal length; OBe, distance Recent studies on morphological variation (and between cochleae; BL, bulla length; CBPL, greatest in some cases of genetic variation) have shown a basial pit length; OL, dentary length; CIBW, Cl tendency for some species of bats that are widely basal breadth; C1C1B, width across C1CI labial distributed in Nusa Tenggara and Maluku surfaces at alveoli; MJM3B, width across M3M3 Tenggara to differentiate morphologically in the labial surfaces at alveoli; C1M3L, maxillary tooth southeastern parts of these provinces, where the row length, from Clalveoli anterior edge to M3 islands in the Inner and Outer Banda Arcs become alveoli posterior edge; M 2L, M 2 crown length; M2W, 214 D.J. Kitchener, A. Suyanto

II 1000l

Figure 1 Locality of Taphozous melanopogon (e) and T. achates (0) used in this study.

M2 crown breadth; SVL, tip of snout to anus length; islands which had both sexes and all age classes TVL, distal tip of tail to anus length; EL, ear length; present (Lombok, Sumbawa, Timor, Roti, Wetar, GTW, greatest tragus breadth; RL, radius length; Kai Besar and Selaru (Tanimbar Group). Graphical D3-SMCL, digit 3-5 metacarpal length; D3Pl-2L, inspection of raw data using residual plots from digit 3 phalanx 1-2 length; TL, tibia length; PES, regression analysis gave no indication of pes length. heteroscedasticity. Because of the number of All skull, dentary and dental characters were interactions tested in this analysis the probability measured with vernier callipers to 0.01 mm and level was set at P

Table 1 Measurements (in mm) for skull, dentary, dental and external body characters of adult Ttphozous a. achates from Wetar Island and T. a. minor subsp. novo from both Yamdena and Kai Besar Islands. X, mean; SD, standard deviation; MIN, minimum; MAX, maximum and N, sample size.

GSL IOW POW CW MW ZW PPL PL DBC BL GBPL DL CIBW

T. a. achates X 21.66 6.35 4.72 10.00 11.38 13.25 10.36 6.24 2.31 5.02 4.01 17.21 1.29 (Wetar Island) SD 0.39 0.23 0.08 0.19 0.17 0.25 0.19 0.19 0.14 0.14 0.14 0.30 0.07 MIN 21.22 5.95 4.65 9.62 11.12 12.88 10.09 6.05 2.09 4.79 3.85 16.82 1.15 MAX 22.55 6.59 4.89 10.25 11.62 13.71 10.67 6.52 2.51 5.13 4.34 17.76 1.37 N 9 9 9 9 9 8 9 9 9 8 9 9 9

T. a. minor X 20.36 6.25 4.57 9.62 11.00 12.76 9.98 5.82 2.12 4.72 3.71 16.32 1.17 subsp. novo SD 0.21 0.29 0.09 0.09 0.10 0.17 0.13 0.17 0.06 0.09 0.11 0.25 0.07 (Yamdena Island) MIN 19.94 5.82 4.48 9.42 10.88 12.42 9.82 5.43 2.04 4.57 3.49 12.96 1.06 MAX 20.61 6.62 4.73 9.78 11.15 13.01 10.21 6.10 2.23 4.80 3.84 16.65 1.28 N 12 12 12 12 12 12 12 12 12 8 12 12 12

T. a. minor X 20.74 6.21 4.65 9.64 11.09 12.97 10.10 6.02 2.15 4.74 3.85 16.56 1.26 subsp. novo SD 0.25 0.23 0.07 0.10 0.09 0.11 0.23 0.14 0.12 0.08 0.12 0.16 0.06 (Kai Besar Island) MIN 20.43 5.91 4.54 9.42 10.96 12.81 9.75 5.77 2.01 4.61 3.56 16.27 1.20 MAX 21.12 6.61 4.81 9.80 11.25 13.19 10.43 6.27 2.41 4.86 4.07 16.80 1.38 N 12 12 12 12 12 12 12 12 12 12 12 12 12 Indonesian Bearded Tomb Bats 215

larger set of characters. This reduced set of island at P

size but the Wetar individuals were generally . variable (F6 88=10.699; P=

Table 1 (continued)

CICIB M 3 M 3 BC1 M 3 L M2L M2W SVL TVL EL GTW RL D3MCL D3PIL D3P2L D4MCL D5MCL TL PES

--,--~-"--,,---"-" -"-,-~~"""""-- . - .._"'~--_._------4.24 9.12 9.35 2.15 2.43 70.6 27.7 21.5 5.1 64.5 57.8 20.3 22.9 46.3 36.7 258 11.6 0.15 0.27 0.22 0.08 0.09 31 2.1 11 0.2 08 11 07 09 11 07 0.6 0.4 3.98 870 9.06 2.07 228 64.9 23.4 19.8 4.9 63.2 555 19.6 211 44.4 355 24.5 11.2 455 9.62 974 2.28 2.61 74.9 30.3 23.1 54 65.4 58.9 21.2 24.2 48.2 374 265 12.2 9 9 9 9 9 9 9 9 9 9 9 9 9 9 9 9 9

3.99 8.77 8.63 1.97 232 69.6 26.8 19.1 47 625 54.9 20.1 21.9 44.4 348 24.3 11.4 007 0.14 0.15 006 007 3.0 1.7 05 0.2 11 08 05 04 1.0 07 04 03 390 859 8.45 1.83 2.14 65.3 240 183 44 602 533 195 21.3 HO 340 23.8 107 409 895 889 206 2.41 75.2 308 197 51 64.1 560 208 22.8 459 359 24.9 119 12 12 12 12 12 12 12 12 12 12 12 12 12 12 12 12 12

4.15 895 8.91 1.96 2.29 735 26.6 203 50 61.2 54.6 19.6 21.2 44.3 343 24.6 114 0.06 0.12 017 0.08 0.08 37 1.3 0.4 0.2 0.8 06 0.4 06 07 06 04 02 403 865 8.63 1.79 2.20 65.2 238 198 47 595 538 19.1 198 43.0 331 239 11.0 429 909 9.12 205 241 797 28.2 20.9 5.2 625 55.5 205 22.1 45.1 35.4 253 11.8 12 12 12 12 12 12 12 12 12 12 12 12 12 12 12 12 12 216 D.J. Kitchener, A. Suyanto

Savu Group Saw Group

/'W~w 0/ 0 D~" j 0 0 -'0 IF EEl Tanimbar Group FWEOFD W ///T-~K \ I \E W K " K/ TT K \, \fl /W '\ T KKKTT K "'\ K // "'T K / ~""''' r...... -"----- K/

A-A-H-C----A ~AG A /lCA A AH AIlGA H I I HG&r HGA I ~AH ~~AA I A AA A HHG C -2 '"AAAA"lICCA C/

wo"C'-, '"<:~ -4 ;;/' A

-6 -4 a b Function 2 Function 2 Figure 2 Canonical variate analysis for both adult male and adult female Bearded Tomb Bats based three island groups (Tanimbar Group; Savu Group, and Lombok Group). The plots of Functions 1 and 2 were based on (a) five selected skull, dentary and dental characters and (b) five selected external characters. The island codes within these three groups were as follows: A, Lombok; B, Alar, C, Timor; D, Roti, E, Savu; F, Semau; G, Mayo; H, Sumbawa; W, Wetar; K, Kai and T, Tanimbar.

Group (Yamdena and Kai Besar)] produced a 75.5% of the variance, separated both the Savu and similar association of these island populations in Tanimbar Groups from the Lombok Group (Figure Discriminant Function space to that produced by 2b). The character that loaded heavily (>0.5) on the above DFA using all islands and 17 characters. Function 1 was tibia length (Table 2b). Function 2, This latter DFA produced two significant which explained 24.5% of the variance, separated Functions (Figure 2a). Function 1 explained 82.5% the Savu and Tanimbar Groups. The character of the variance and Function 2, 17.5%. All loading heavily (>0.05) on Function 2 was digit 3 individuals were correctly classified to their metacarpal length (Table 2b). A total of 87.9% of appropriate island population. individuals were classified to their correct island The Lombok Group separated from both the group. Misclassifications were as follows: Two Savu and Tanimbar Groups on Function 1. The Savu Group animals to the Tanimbar Group; five characters loading heavily (>0.5) on Function 1 Lombok Group animals to the Savu Group and 10 were dentary length and greatest basial pit length Lombok Group animals to the Tanimbar Group. (Table 2a). The Tanimbar Group separated from The Lombok, Savu and Tanimbar Groups were the Savu Group and partially from the Lombok clearly separated in Discriminant Function space Group on Function 2. The characters loading based on skull, dentary and dental characters, but heavily (>0.5) on Function 2 were greatest basial less clearly so on external characters. Further, it is pit length and M2 length (Table 2). apparent from these plots that within these Groups there is little morphological differentiation between External characters the islands. This is indicated by the observation All external characters, except snout to vent that the Lombok Group cluster is defined in DF length (which showed an interaction between sex space by individuals from a single island (Lombok) and island), were included in a DFA of these and similarly the Savu Group is defined by Wetar characters based on the same three groups individuals. The Yamdena and Kai Besar identified above (Lombok, Savu, and Tanimbar). specimens also overlap considerably (Figure 2). In This DFA was also run using a reduced set of five the previous study by Kitchener et al. (1993a) the characters (tibia length, digit 3 metacarpal length, Lombok Group represent Tapho2ouS m. melanopogon digit 3 phalanx 1 length, digit 5 metacarpal length and the Savu Group T. achates. Consequently the and ear length) and it this latter DFA that is Wetar Island individuals classify as T. achates. The presented here. The DFA extracted two very Tanimbar Group individuals are morphologically significant Functions. Function 1, which explained distinct from both these above taxa. However, the Indonesian Bearded Tomb Bats 217

Table 2 Standardised and unstandardised (in brackets) Kepulauan Kai (05°39'5, 132°59'E), mist netted from Canonical Variate Function coefficients for three the mouth of an open well-lit cave on the beach, by island groupings (the Lombok, Savu and D.J. Kitchener and R.A. How on 19 October 1992. Tanimbar groups see text) using a reduced set of five (a) skull, dentary and dental characters and (b) external characters. Paratypes Table 2a Yamdena Island, Kepulauan Tanimbar Desa Olilit, 1 km 5 5aumlaki (from a World War II Character Function 1 Function 2 Japanese made tunnel) (7°59'5, 131 °22'E), 2 00 9 9 Dentary length 0.9925 (3.9673) -00351 (-0.1404) 9, WAM M (43527, 43529-31, 43538, 43541-43, Basial pit length -0.6740 (-5.1426) 0.5109 (3.8982) 43547-49). 2 km 5 DesaLat Dalam, (from a large Bulla length 04514 (-35766) 04798 (3.8009) 'doline' cave, among a number of other limestone M2 length 0.1256 (2.()740) 06495 (10.7288) caves) (7°03'5, 131°0TE), 1 0, WAM M 43672. Interorbital breadth 03673 (19550) -01179 (-0.6275) Kai Besar Island, Kepulauan Kai, beach cave, Constant -42.9653 -5Cl.0386 Pulau (= islet) Kelapa, 2 km W Elat, Kai Besar Variance Island (05°39'5, 132°59'E), 4 00 8 9 9, WAM M explained (%) 82.5 17.5 (42886-88, 42890-91, 42893-94, 42896-97, 42899­ ._-_ __._------900,42902). Table 2b Character Function 1 Function 2 Diagnosis Tapho2ollS achates minor differs from T. a. acltates Tibia length 0.8558 (0.8429) 0.1265 (0.1246) in averaging smaller in all characters measured Digit 3 metacarpal length -0.2202 (-0.1719) 0.7704 (0.6014) (Table 1 and Kitchener et al. 1993: table 2)). For Ear length 0.4127 (0.3279) 0.2918 (0.2318) example, greatest skull length 20.55 (19.94-21.12) Digit 3 phalanx 1 24 v. 21.54 (20.91-22.55) 28; interorbital breadth length -04681 (-0.6943) -0.0529 (-0.0785) 6.23 (5.82-6.62) 24 V. 6.48 (5.95-6.95) 28; zygomatic Digit 5 metacarpal width 12.87 (12.42-13.19) 24 V. 13.22 (12.88-13.78) length -0.1177 (-0.0836) 0.3315 (0.2496) 26; bulla length 4.73 (4.57-4.86) 20 V. 4.97 (4.75­ Constant 04272 -48.8648 5.55) 27; greatest basial pit length 3.78 (3.49-4.07) Variance 24 v. 4.04 (3.85-4.34) 28; l M3 (alveoli) length 8.77 explained (%) 75.5 24.5 e (8.45-9.12) 24 V. 9.23 (8.85-9.74) 28; M2 (crown) length 1.96 (1.79-2.06) 24 V. 2.15 (2.07-2.28) 28; tail to vent length 26.7 (23.8-30.8) 24 V. 27.3 (23.4-31.6) DFA of all 11 islands and all skull, dentary and 28; radius length 61.8 (59.5-64.1) 24 V. 63.8 (61.1­ dental characters indicate that on Function 3 65.4) 28; tibia length 24.5 (23.8-25.3) 24 V. 25.5 individuals from the 5avu and Tanimbar Groups (23.3-26.8) 28. Also M2 crown length and bulla overlap considerably (graph not presented) length shorter relative to least interorbital breadth indicating that individuals from the Tanimbar (Figure 3a,b); dentary length generally shorter Group were associated in OF space with T. achates relative to greatest basial pit length (Figure 4); and rather that with T. m. melanopogon. In the following digit 3 metacarpal length shorter relative to tibia section the populations from Tanimbar and Kai length (Figure 5). Islands are described as a new subspecies of T. It differs from Taplto2ous m. melanopogon from achates. these southern Indonesian islands by having mature adult males with a pale brown rather than a black throat beard. Most skull, dentary and TAXONOMY dental characters average smaller. For example cranial breadth 9.63 (9.42-9.80) 24 V. 9.78 (9.46­ Tapho20llS achates minor subsp. novo Kitchener 10.22) 92; distance between cochleae 2.13 (2.01­ 2.41) 24 V. 2.27 (1.89-2.66) 92; bulla length 4.73 Holotype (4.57-4.86) 20 v. 4.97 (1.59-5.42) 91; greatest basial Museum Zoologicum Bogoriense (MZB) No. pit length 3.78 (3.49-4.07) 24 v. 4.09 (3.73-4.36) 92; 15908 (field number M42899), adult male, carcase radius length 61.8 (59.5-64.1) 24 v. 63.4 (60.9-66.1) fixed in 10')';, formalin and preserved in 70% 90. 50me characters average larger. For example, ethanol, skull removed, cleaned and separate, liver least interorbital breadth 6.23 (5.82-6.62) 24 v. 5.99 stored in ultrafreeze at the Western Australian (5.60-6.33) 92; zygomatic width 12.87 (12.42-13.19) Museum. 24 V. 12.60 (11.98-13.16) 92; tail to vent length 26.7 (23.8-30.8) 24 v. 24.6 (18.8-29.5) 90; and tibia length Type locality 24.5 (23.8-25.3) 24 V. 23.3 (19.9-25.5) 90. Least Pulau Kelapa, 2 km W Elat, Kai Besar Island interorbital breadth greater relative to M2 length 218 D.J. Kitchener, A. Suyanto

2.3 T:o. achates o o Tm. melanopogon o 2.2

T.a. adwles

5.2 A F o .c H 2.1 00 \A W...____W C ~ W\---- A A"':. ,,':<~YW .c !'! HH~ A~I 00 :; I> c c:l H CH\'I/CG A AB / ~ T.a minor 5,0 GACAA A AA 0 E ~ AGA CHHCH A ~HAAAB// OFP / HA W GGC \ 0 F AA AA A\F HA)?piA A W 1.9 AK ~ Cl \ 0 AAA/_~ 4.8 / G TA T-K-F\T~0 A AK T ~ H ~- KK \E-d'E

1.8 \ K K I ~/ TKK J T.a minor 4.6 ~------

56 5.8 6.0 6.2 6.4 6.6 6.8 70 5.6 5.8 6.0 6·2 6 .• 6.8 7.0 a b Interorbital breadth Interorbital breadth Figure 3 Plots of least interorbital breadth versus (a) M2 length and (b) bulla length for both adult male and adult female T. m. melanopogon, T. a. achaies and T. a. minor for all island populations. Codes for islands are as for Figure 2.

and bulla length (Figure 3a,b); greatest basial pit populations of T. a. achates were included within length shorter relative to dentary length (Figure 4) the Wetar Island population cluster in discriminant and tibia length generally longer relative to digit 3 function space. This suggests that the Rotinese metacarpal length (Figure 5). population had not noticeably differentiated from the nominate subspecies as suggested by Kitchener Description et al. (1993a). Apart from the differences mentioned in the Interestingly, T. achates occurred on Tanimbar above diagnoses T. a. minor is very similar morphologically and in its pelage colour to T. a. W, T.a. achater achates. 17.6 I~~O-O\ Distribution ..,rOD ~ E/W Known only from Yamdena Island and Kai Besar, 11.2 Maluku Tenggara, eastern Indonesia. / WF EO F/ W-F-F\WW / E "E_O Etymology 16.8 K---W-K K The name minor is from the latin referring to the H/~H", T.amin'" T/ : K F/ TKKKK / HAB A small size of the Yamdena and Kai Besar form of T. :A--/~ / GH'ICHA Cl achates compared to members of the nominate 16.4 K K T./"I~ H hA G A Au4 TT / /. HA H CH 't TK AHG G C GA subspecies. /A AH GA AAH / T / AA HG H ; T A A A/ "'-. A AHC AB 18,0 "'-T__ AT AA AG AA \, AA AACA HA A/A DISCUSSION C-A "" /GA.... 1:111. melanopogon The Bearded Tomb Bats from Tanimbar and Kai 15.6 ""G'A Islands are most similar to T. a. achates but differ in

being generally smaller in most characters; some 3.4 3.6 3.8 4.0 4.2 4.4 skull and external characters also differ in shape. For example, M2 crown length, bulla length and Basial pit length digit 3 metacarpal length. Specimens from Wetar Figure 4 Plots of dentary length versus greatest basial Island were T. a. achates and were not pit length for both adult male and adult differentiated morphologically from the other female T. m. melanopogon, T. a. achates and T. known populations of this nominate subspecies on a, minor for all island populations. Codes for Semau, Roti, and Savu Islands. In fact these latter islands are as for Figure 2. Indonesian Bearded Tomb Bats 219

27 T(L D throughout Indonesia, also differentiates into a dis­ tinct form on Wetar and Islands (Kitchener, o IN //OFF unpublished data). To minor OW FIN WED ww A E K EDD o T~ /~ D Other specimens examined .r 1 K ,A--WAG GAH K G KT C AH 24~ A- KA E A AA AG TllpllOZOllS achlltes llclllltcS T- T T~K-T H A HA A A A HA H Desa Ipokil, Wetar Island, Nusa Tenggara Timur HHH A FC C H A A GC (7°5CrS, 126°16'E), 4 0 (~ 5 22 WAM M(44510, G HA A A A AB C CH A H 44520-27). H A A A A 22 CA C A A A A ACKNOWLEDGEMENTS " A We are grateful for the support of Mr A. Reeves, Director, Western Australian Museum and to Or Soetikno, Director, Puslitbang Biologi, and Mr M. Amir, Director, Balitbang Zoologi, Bogor Indonesia 53 54 55 56 57 56 59 60 and the Directors of Sub Balai Konservasi in Nusa Tenggara Barat (Ir. P. Supriadi), Nusa Tenggara Digit 3 metacarpal length Timur (Ir. J. Mochtar) and Maluku Tenggara (Ir. J. Rustandi). Figure 5 Plot of tibia length versus digit 3 metacarpal length for both adult male and adult female A number of colleagues assisted in the field work T. m. melanopogon, T. a. achates and T. a. minor of whom Or R.A. How, Mr R.E. Johnstone, for all island populations. Codes for islands Western Australian Museum, and Ir. are as for Figure 2. Maharadatunkamsi and Ir. 1. Maryanto, Balitbang Zoologi, were principal. Mr C. Keast, Western Australian Museum, assisted in the preparation of and Kai Islands as predicted by Kitchener et al. skulls for this study. Mrs S. Dalton, Western (1993a). Its distribution on the islands to the Australian Museum, typed the manuscript. immediate south and north of Timor islands add The field work was supported by a number of credence to its presumed presence also on Timor grants, including National Geographic Society Island (see Kitchener et al. 1993a). We have not Research Grants, Washington and the Australian examined the specimen from Kai Island which Nature Conservation Agency, Canberra. Freight Koopman and Gordon (1992) attributed to T. costs were subsidised by Garuda Airlines. melanopogon, but consider that it most probably also represents T. achates minor. The morphological differentiation of T. achates on REFERENCES the eastern islands of Kai and Tanimbar reflects a Kitchener, D.J., How, R.A., Cooper, N. and Suyanto, A. pattern of intraspecific differentiation observed in (1992). Hipposidcros diadema (Chiroptera, other mammal species in the region. For example, Hipposideridae) in the , Hipposideros sumbae, Pteropus lombocensis and Indonesia: Taxonomony and geographic Cynopterlls nusatenggara, all Nusa Tenggara morphological variation. R.ecords of the Western endemics, differentiate morphologically into Australian Museum 16: 1-60. western and eastern forms (Kitchener and Kitchener, D.J. and Maryanto, I. (1993). Taxonomic re­ Maryanto 1993, Kitchener et al. 1995c,d). The bats appraisal of the Hipposideros larvatus species complex Hipposideros diadema, Nyctimene albiventer, N. (Chiroptera: Hipposideridae) in the Greater Sunda cephalotes, Syconycteris allstralis, Myotis adversus, Is, Indonesia. Records of the Western Australian Rhinolophlls affinis and R. simplex, all species with Museum 16: 119-173. more widespread distributions, also differentiate Kitchener, D.J. and Schmitt, L.H., Hisheh, S., How, R.A., morphologically within this region (Kitchener et al Cooper, N. and Maharadatunkamsi (1993a). Morphological and genetic variation in the Bearded 1992, Kitchener et al. 1993; Kitchener et al.; in press Tomb Bats (Taphozous: Emballonuridae) of Nusa 1995a-d; Maharadatunkamsi 1990). This Tenggara, Indonesia. Mammalia 57: 63-83. differentiation becomes more extreme on the more Kitchener, D.J., Packer, W.e. and Maryanto, I (1993b). remote eastern islands of the Inner Arc, or on the Taxonomic status of Nyctimene (ChiroptelT islands of the Outer Banda Arc that have been Pteropodidae) from the Banda.. Kai and Aru Islands, continuously isolated by seas from other island MaIuku, Indonesia implications for biogeography populations. Interestingly, the House Mouse, Mus Records of tile Western Australian Museum 16: 399--417 musclllus castanells, which is widely distributed Kitchener, D.J., Schmitt, L.H., StranD, P, Wheeler, A. 220 D.J. Kitchener, A. Suyanto

and Suyanto, A. (1995a). Taxonomy of Rhinolophus Western Australian Museum 17: 125-142. simplex Anderson, 1905 (Chiroptera: Rhinolophidae) Kitchener, DJ, Packer, W.e. and Maryanto, 1. (in press). in Nusa Tenggara and Maluku, Indonesia, Records of Morphological variation in Maluku populations of the Western Australian Museum 17: 191-212. Syconycteris australis (Peters, 1867) (Chiroptera: Kitchener, D.J., Cooper, N. and Maryanto, 1. (in press, Pteropodidae). Records of the Western Australian b). The Myotis adversus (Chiroptera, Vespertilionidae) Museum. species complex in Eastern Indonesia, Australia, Koopman, K.F. and Gordon, L.K. (1992). Systematic and Solomon Islands. Records of notes on a collection of bats from the Western Australian Museum. (Indonesia: Moluccas). American Museum Novitates Kitchener, D.J., Packer, W.e. and Maharadatunkamsi 3035: 1-8. (1995c). Morphological variation in Pteropus Maharadatunkarnsi, (1990). Morphological and genetic lombocensis (Chiroptera: Pteropodidae) in Nusa variation in Rhinolophus affinis from Jawa and Nusa Tenggara, Indonesia. Records ofthe Western Australian Tenggara, Indonesia. Postgraduate Diploma of Museum 17: 61-67. Science Thesis, University of Western Australia. Kitchener, D.J., Packer, W.e. and Suyanto, A. (1995d). Systematic review of Nyctimene cephalotes and N. albiventer (Chiroptera: Pteropodidae) in the Maluku Manuscript received 25 May 1994; accepted 13 February and regions, Indonesia. Records of the 1995.